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Linked References

1. A. Yoshimura, T. Naka, and M. Kubo, “SOCS proteins, cytokine signalling and immune regulation,” Nature Reviews Immunology, vol. 7, no. 6, pp. 454–465, 2007. View at Publisher · View at Google Scholar · View at PubMed · View at Scopus
2. J. Piessevaux, D. Lavens, F. Peelman, and J. Tavernier, “The many faces of the SOCS box,” Cytokine and Growth Factor Reviews, vol. 19, no. 5-6, pp. 371–381, 2008. View at Publisher · View at Google Scholar · View at PubMed · View at Scopus
3. L. N. Akhtar and E. N. Benveniste, “Viral exploitation of host SOCS protein functions,” Journal of Virology, vol. 85, no. 5, pp. 1912–1921, 2011. View at Publisher · View at Google Scholar · View at PubMed
4. C. J. Secombes, T. Wang, and S. Bird, “The interleukins of fish,” Developmental and Comparative Immunology. In press. View at Publisher · View at Google Scholar · View at PubMed
5. T. Wang, W. Huang, M. M. Costa, and C. J. Secombes, “The γ-chain cytokine/receptor system in fish: more ligands and receptors,” Fish and Shellfish Immunology, vol. 31, pp. 673–687, 2001. View at Publisher · View at Google Scholar · View at PubMed
6. S. Hong, J. Zou, M. Crampe et al., “The production and bioactivity of rainbow trout (Oncorhynchus mykiss) recombinant IL-1β,” Veterinary Immunology and Immunopathology, vol. 81, no. 1-2, pp. 1–14, 2001. View at Publisher · View at Google Scholar · View at Scopus
7. P. Díaz-Rosales, S. Bird, T. Wang et al., “Rainbow trout interleukin-2: cloning, expression and bioactivity analysis,” Fish and Shellfish Immunology, vol. 27, no. 3, pp. 414–422, 2009. View at Publisher · View at Google Scholar · View at PubMed · View at Scopus
8. M. M. Costa, T. Maehr, P. Diaz-Rosales, C. J. Secombes, and T. Wang, “Bioactivity studies of rainbow trout (Oncorhynchus mykiss) interleukin-6: effects on macrophage growth and antimicrobial peptide gene expression,” Molecular Immunology, vol. 31, no. 1, pp. 107–117, 2011. View at Publisher · View at Google Scholar · View at PubMed
9. N. O. Harun, J. Zou, Y. A. Zhang, P. Nie, and C. J. Secombes, “The biological effects of rainbow trout (Oncorhynchus mykiss) recombinant interleukin-8,” Developmental and Comparative Immunology, vol. 32, no. 6, pp. 673–681, 2008. View at Publisher · View at Google Scholar · View at PubMed · View at Scopus
10. L. Grayfer, J. W. Hodgkinson, S. J. Hitchen, and M. Belosevic, “Characterization and functional analysis of goldfish (Carassius auratus L.) interleukin-10,” Molecular Immunology, vol. 48, no. 4, pp. 563–571, 2011. View at Publisher · View at Google Scholar · View at PubMed · View at Scopus
11. T. Wang, J. W. Holland, A. Carrington, J. Zou, and C. J. Secombes, “Molecular and functional characterization of IL-15 in rainbow trout Oncorhynchus mykiss: a potent inducer of IFN-γ expression in spleen leukocytes,” Journal of Immunology, vol. 179, no. 3, pp. 1475–1488, 2007. View at Scopus
12. T. Wang, P. Diaz-Rosales, M. M. Costa, et al., “Functional characterization of a nonmammalian IL-21: rainbow trout Oncorhynchus mykiss IL-21 upregulates the expression of the Th cell signature cytokines IFN-gamma, IL-10, and IL-22,” Journal of Immunology, vol. 186, no. 2, pp. 708–721, 2011.
13. M. M. Monte, J. Zou, T. Wang, A. Carrington, and C. J. Secombes, “Cloning, expression analysis and bioactivity studies of rainbow trout (Oncorhynchus mykiss) interleukin-22,” Cytokine, vol. 55, no. 1, pp. 62–73, 2011. View at Publisher · View at Google Scholar · View at PubMed
14. J. Zou, S. Peddie, G. Scapigliati et al., “Functional characterisation of the recombinant tumor necrosis factors in rainbow trout, Oncorhynchus mykiss,” Developmental and Comparative Immunology, vol. 27, no. 9, pp. 813–822, 2003. View at Publisher · View at Google Scholar · View at Scopus
15. M. C. Ordás, M. M. Costa, F. J. Roca et al., “Turbot TNFα gene: molecular characterization and biological activity of the recombinant protein,” Molecular Immunology, vol. 44, no. 4, pp. 389–400, 2007. View at Publisher · View at Google Scholar · View at PubMed · View at Scopus
16. J. Zou, C. Tafalla, J. Truckle, and C. J. Secombes, “Identification of a second group of type I IFNs in fish sheds light on IFN evolution in vertebrates,” Journal of Immunology, vol. 179, no. 6, pp. 3859–3871, 2007. View at Scopus
17. J. Zou, A. Carrington, B. Collet et al., “Identification and bioactivities of IFN-γ in rainbow trout Oncorhynchus mykiss: the first Th1-type cytokine characterized functionally in fish,” Journal of Immunology, vol. 175, no. 4, pp. 2484–2494, 2005. View at Scopus
18. J. A. Arts, E. J. Tijhaar, M. Chadzinska, H. F. Savelkoul, and B. M. Verburg-van Kemenade, “Functional analysis of carp interferon-γ: evolutionary conservation of classical phagocyte activation,” Fish and Shellfish Immunology, vol. 29, no. 5, pp. 793–802, 2010. View at Publisher · View at Google Scholar · View at PubMed · View at Scopus
19. Y. Nakatani, H. Takeda, Y. Kohara, and S. Morishita, “Reconstruction of the vertebrate ancestral genome reveals dynamic genome reorganization in early vertebrates,” Genome Research, vol. 17, no. 9, pp. 1254–1265, 2007. View at Publisher · View at Google Scholar · View at PubMed · View at Scopus
20. H. J. Jin, J. Z. Shao, L. X. Xiang, H. Wang, and L. L. Sun, “Global identification and comparative analysis of SOCS genes in fish: insights into the molecular evolution of SOCS family,” Molecular Immunology, vol. 45, no. 5, pp. 1258–1268, 2008. View at Publisher · View at Google Scholar · View at PubMed · View at Scopus
21. T. Wang, Q. Gao, P. Nie, and C. J. Secombes, “Identification of suppressor of cytokine signalling (SOCS) 6, 7, 9 and CISH in rainbow trout Oncorhynchus mykiss and analysis of their expression in relation to other known trout SOCS,” Fish and Shellfish Immunology, vol. 29, no. 4, pp. 656–667, 2010. View at Publisher · View at Google Scholar · View at PubMed · View at Scopus
22. S. E. Altschul, W. Gish, W. Miller, E. W. Myers, and D. J. Lipman, “Basic local alignment search tool,” Journal of Molecular Biology, vol. 215, no. 3, pp. 403–410, 1990. View at Publisher · View at Google Scholar · View at PubMed · View at Scopus
23. T. Wang and C. J. Secombes, “Complete sequencing and expression of three complement components, C1r, C4 and C1 inhibitor, of the classical activation pathway of the complement system in rainbow trout Oncorhynchus mykiss,” Immunogenetics, vol. 55, no. 9, pp. 615–628, 2003. View at Publisher · View at Google Scholar · View at PubMed · View at Scopus
24. T. Wang, P. C. Hanington, M. Belosevic, and C. J. Secombes, “Two macrophage colony-stimulating factor genes exist in fish that differ in gene organization and are differentially expressed,” Journal of Immunology, vol. 181, no. 5, pp. 3310–3322, 2008. View at Scopus
25. J. J. Campanella, L. Bitincka, and J. Smalley, “MatGAT: an application that generates similarity/identity matrices using protein or DNA sequences,” BMC Bioinformatics, vol. 4, article 29, 2003. View at Publisher · View at Google Scholar · View at PubMed
26. B. A. Croker, H. Kiu, and S. E. Nicholson, “SOCS regulation of the JAK/STAT signalling pathway,” Cytokine & Growth Factor Reviews, vol. 19, no. 4, pp. 371–381, 2008.
27. L. W. Waiboci, C. M. Ahmed, M. G. Mujtaba et al., “Both the suppressor of cytokine signaling 1 (SOCS-1) kinase inhibitory region and SOCS-1 mimetic bind to JAK2 autophosphorylation site: implications for the development of a SOCS-1 antagonist,” Journal of Immunology, vol. 178, no. 8, pp. 5058–5068, 2007. View at Scopus
28. A. N. Bullock, J. E. Debreczeni, A. M. Edwards, M. Sundström, and S. Knapp, “Crystal structure of the SOCS2-elongin C-elongin B complex defines a prototypical SOCS box ubiquitin ligase,” Proceedings of the National Academy of Sciences of the United States of America, vol. 103, no. 20, pp. 7637–7642, 2006. View at Publisher · View at Google Scholar · View at PubMed · View at Scopus
29. R. Chenna, H. Sugawara, T. Koike et al., “Multiple sequence alignment with the Clustal series of programs,” Nucleic Acids Research, vol. 31, no. 13, pp. 3497–3500, 2003. View at Publisher · View at Google Scholar · View at Scopus
30. M. Rechsteiner and S. W. Rogers, “PEST sequences and regulation by proteolysis,” Trends in Biochemical Sciences, vol. 21, no. 7, pp. 267–271, 1996. View at Publisher · View at Google Scholar · View at Scopus
31. J. J. Babon, S. Yao, D. P. DeSouza et al., “Secondary structure assignment of mouse SOCS3 by NMR defines the domain boundaries and identifies an unstructured insertion in the SH2 domain,” FEBS Journal, vol. 272, no. 23, pp. 6120–6130, 2005. View at Publisher · View at Google Scholar · View at PubMed · View at Scopus
32. K. Tamura, D. Peterson, N. Peterson, G. Stecher, M. Nei, and S. Kumar, “MEGA5: molecular evolutionary genetics analysis using maximum likelihood, evolutionary distance, and maximum parsimony methods,” Molecular Biology and Evolution, vol. 28, no. 10, pp. 2731–2739, 2011.
33. M. Kozak, “Pushing the limits of the scanning mechanism for initiation of translation,” Gene, vol. 299, no. 1-2, pp. 1–34, 2002. View at Publisher · View at Google Scholar
34. M. Muffato, A. Louis, C. E. Poisnel, and H. R. Crollius, “Genomicus: a database and a browser to study gene synteny in modern and ancestral genomes,” Bioinformatics, vol. 26, no. 8, pp. 1119–1121, 2010. View at Publisher · View at Google Scholar · View at PubMed · View at Scopus
35. T. Wang and C. J. Secombes, “Rainbow trout suppressor of cytokine signalling (SOCS)-1, 2 and 3: molecular identification, expression and modulation,” Molecular Immunology, vol. 45, no. 5, pp. 1449–1457, 2008. View at Publisher · View at Google Scholar · View at PubMed · View at Scopus
36. B. J. Baker, L. N. Akhtar, and E. N. Benveniste, “SOCS1 and SOCS3 in the control of CNS immunity,” Trends in Immunology, vol. 30, no. 8, pp. 392–400, 2009. View at Publisher · View at Google Scholar · View at PubMed · View at Scopus
37. H. J. Jin, L. X. Xiang, and J. Z. Shao, “Identification and characterization of suppressor of cytokine signaling 1 (SOCS-1) homologues in teleost fish,” Immunogenetics, vol. 59, no. 8, pp. 673–686, 2007. View at Publisher · View at Google Scholar · View at PubMed · View at Scopus
38. H. J. Jin, J. Z. Shao, and L. X. Xiang, “Identification and characterization of suppressor of cytokine signaling 3 (SOCS-3) homologues in teleost fish,” Molecular Immunology, vol. 44, no. 5, pp. 1042–1051, 2007. View at Publisher · View at Google Scholar · View at PubMed · View at Scopus
39. Z. G. Xiao, H. Liu, J. P. Fu, W. Hu, Y. P. Wang, and Q. L. Guo, “Cloning of common carp SOCS-3 gene and its expression during embryogenesis, GH-transgene and viral infection,” Fish and Shellfish Immunology, vol. 28, no. 2, pp. 362–371, 2010. View at Publisher · View at Google Scholar · View at PubMed · View at Scopus
40. M. Zhang, Z. Z. Xiao, and L. Sun, “Suppressor of cytokine signaling 3 inhibits head kidney macrophage activation and cytokine expression in Scophthalmus maximus,” Developmental and Comparative Immunology, vol. 35, no. 2, pp. 174–181, 2011. View at Publisher · View at Google Scholar · View at PubMed · View at Scopus
41. I. D. Dimitriou, L. Clemenza, A. J. Scotter, G. Chen, F. M. Guerra, and R. Rottapel, “Putting out the fire: coordinated suppression of the innate and adaptive immune systems by SOCS1 and SOCS3 proteins,” Immunological Reviews, vol. 224, no. 1, pp. 265–283, 2008. View at Publisher · View at Google Scholar · View at PubMed · View at Scopus
42. E. Kario, M. D. Marmor, K. Adamsky et al., “Suppressors of cytokine signaling 4 and 5 regulate epidermal growth factor receptor signaling,” Journal of Biological Chemistry, vol. 280, no. 8, pp. 7038–7048, 2005. View at Publisher · View at Google Scholar · View at PubMed · View at Scopus
43. L. A. O'Sullivan, S. M. Noor, M. C. Trengove et al., “Suppressor of cytokine signaling 1 regulates embryonic myelopoiesis independently of its effects on T cell development,” Journal of Immunology, vol. 186, no. 8, pp. 4751–4761, 2011. View at Publisher · View at Google Scholar · View at PubMed
44. M. S. Kim and K. H. Kim, “Inhibition of viral hemorrhagic septicemia virus replication using a short hairpin RNA targeting the G gene,” Archives of Virology, vol. 156, no. 3, pp. 457–464, 2011. View at Publisher · View at Google Scholar · View at PubMed · View at Scopus
45. C. Tafalla, J. Coll, and C. J. Secombes, “Expression of genes related to the early immune response in rainbow trout (Oncorhynchus mykiss) after viral haemorrhagic septicemia virus (VHSV) infection,” Developmental and Comparative Immunology, vol. 29, no. 7, pp. 615–626, 2005. View at Publisher · View at Google Scholar · View at PubMed · View at Scopus
46. D. M. Stone, H. W. Ferguson, P. A. Tyson et al., “The first report of viral haemorrhagic septicaemia in farmed rainbow trout, Oncorhynchus mykiss (Walbaum), in the United Kingdom,” Journal of Fish Diseases, vol. 31, no. 10, pp. 775–784, 2008. View at Publisher · View at Google Scholar · View at PubMed · View at Scopus
47. R. P. Hedrick, T. S. McDowell, M. Gay, G. D. Marty, M. P. Georgiadis, and E. MacConnell, “Comparative susceptibility of rainbow trout Oncorhynchus mykiss and brown trout Salmo trutta to Myxobolus cerebralis, the cause of salmonid whirling disease,” Diseases of Aquatic Organisms, vol. 37, no. 3, pp. 173–183, 1999. View at Scopus
48. S. Hadidi, G. W. Glenney, T. J. Welch, J. T. Silverstein, and G. D. Wiens, “Spleen size predicts resistance of rainbow trout to Flavobacterium psychrophilum challenge,” Journal of Immunology, vol. 180, no. 6, pp. 4156–4165, 2008. View at Scopus
49. S. J. Haig, R. L. Davies, T. J. Welch, R. A. Reese, and D. W. Verner-Jeffreys, “Comparative susceptibility of Atlantic salmon and rainbow trout to Yersinia ruckeri: relationship to O antigen serotype and resistance to serum killing,” Veterinary Microbiology, vol. 147, no. 1-2, pp. 155–161, 2011. View at Publisher · View at Google Scholar · View at PubMed
50. C. R. W. Gould, Studies on proliferative kidney disease (PKD) in rainbow trout, Oncorhynchus mykiss, Ph.D. thesis, University of Aberdeen, 1995.
51. J. L. Ho and J. R. Lapa e Silva, “Promotion of a down-modulated lung immune state may be a strategy by M. tuberculosis to foster active disease and persistence,” Discovery Medicine, vol. 9, no. 44, pp. 34–41, 2010.
52. E. Tobback, A. Decostere, K. Hermans, F. Haesebrouck, and K. Chiers, “Yersinia ruckeri infections in salmonid fish,” Journal of Fish Diseases, vol. 30, no. 5, pp. 257–268, 2007. View at Publisher · View at Google Scholar · View at PubMed
53. S. Bertholet, H. L. Dickensheets, F. Sheikh, A. A. Gam, R. P. Donnelly, and R. T. Kenney, “Leishmania donovani-induced expression of suppressor of cytokine signaling 3 in human macrophages: a novel mechanism for intracellular parasite suppression of activation,” Infection and Immunity, vol. 71, no. 4, pp. 2095–2101, 2003. View at Publisher · View at Google Scholar
54. S. Zimmermann, P. J. Murray, K. Heeg, and A. H. Dalpke, “Induction of suppressor of cytokine signaling-1 by Toxoplasma gondii contributes to immune evasion in macrophages by blocking IFN-γ signaling,” Journal of Immunology, vol. 176, no. 3, pp. 1840–1847, 2006.
55. R. P. Hedrick, E. MacConnell, and P. de Kinkelin, “Proliferative kidney disease of salmonid fish,” Annual Review of Fish Diseases, vol. 3, no. C, pp. 277–290, 1993.
56. D. S. Grabner and M. El-Matbouli, “Transmission of Tetracapsuloides bryosalmonae (Myxozoa: Malacosporea) to Fredericella sultana (Bryozoa: Phylactolaemata) by various fish species,” Diseases of Aquatic Organisms, vol. 79, no. 2, pp. 133–139, 2008. View at Publisher · View at Google Scholar
57. E. U. Canning, A. Curry, S. W. Feist, M. Longshaw, and B. Okamura, “Tetracapsula bryosalmonae n.sp. for PKX organism, the cause of PKD in Salmonid fish,” Bulletin of the European Association of Fish Pathologists, vol. 19, no. 5, pp. 203–206, 1999.
58. J. W. Holland, B. Okamura, H. Hartikainen, and C. J. Secombes, “A novel minicollagen gene links cnidarians and myxozoans,” Proceedings of the Royal Society B, vol. 278, no. 1705, pp. 546–553, 2011. View at Publisher · View at Google Scholar · View at PubMed
59. S. W. Feist, M. Longshaw, E. U. Canning, and B. Okamura, “Induction of proliferative kidney disease (PKD) in rainbow trout Oncorhynchus mykiss via the bryozoan Fredericella sultana infected with Tetracapsula bryosalmonae,” Diseases of Aquatic Organisms, vol. 45, no. 1, pp. 61–68, 2001.
60. D. J. Morris and A. Adams, “Transmission of Tetracapsuloides bryosalmonae (Myxozoa: Malacosporea), the causative organism of salmonid proliferative kidney disease, to the freshwater bryozoan Fredericella sultana,” Parasitology, vol. 133, no. 6, pp. 701–709, 2006. View at Publisher · View at Google Scholar · View at PubMed
61. H. W. Ferguson and E. A. Needham, “Proliferative kidney disease in rainbow trout Salmo gairdneri Richardson,” Journal of Fish Diseases, vol. 1, pp. 91–108, 1978.
62. R. S. Clifton-Hadley, D. Bucke, and R. H. Richards, “A study of the sequential clinical and pathological changes during proliferative kidney disease in rainbow trout, Salmo gairdneri Richardson,” Journal of Fish Diseases, vol. 10, pp. 335–352, 1987.
63. T. Wang, J. W. Holland, S. A. Martin, and C. J. Secombes, “Sequence and expression analysis of two T helper master transcription factors, T-bet and GATA3, in rainbow trout Oncorhynchus mykiss and analysis of their expression during bacterial and parasitic infection,” Fish and Shellfish Immunology, vol. 29, no. 5, pp. 705–715, 2010. View at Publisher · View at Google Scholar · View at PubMed · View at Scopus
64. R. G. Rosenfeld and V. Hwa, “The growth hormone cascade and its role in mammalian growth,” Hormone Research, vol. 71, no. 2, pp. 36–40, 2009. View at Publisher · View at Google Scholar · View at PubMed · View at Scopus
65. A. L. Studzinski, D. V. Almeida, C. F. C. Lanes, M. D. Figueiredo, and L. F. Marins, “SOCS1 and SOCS3 are the main negative modulators of the somatotrophic axis in liver of homozygous GH-transgenic zebrafish (Danio rerio),” General and Comparative Endocrinology, vol. 161, no. 1, pp. 67–72, 2009. View at Publisher · View at Google Scholar · View at PubMed · View at Scopus
66. M. F. Flajnik and M. Kasahara, “Origin and evolution of the adaptive immune system: genetic events and selective pressures,” Nature Reviews Genetics, vol. 11, no. 1, pp. 47–59, 2010. View at Publisher · View at Google Scholar · View at PubMed · View at Scopus
67. K. S. Kassahn, V. T. Dang, S. J. Wilkins, A. C. Perkins, and M. A. Ragan, “Evolution of gene function and regulatory control after whole-genome duplication: comparative analyses in vertebrates,” Genome Research, vol. 19, no. 8, pp. 1404–1418, 2009. View at Publisher · View at Google Scholar · View at PubMed · View at Scopus
68. J. H. Postlethwait, “The zebrafish genome in context: ohnologs gone missing,” Experimental Zoology Part B: Molecular and Developmental Evolution, vol. 308, no. 5, pp. 563–577, 2007. View at Publisher · View at Google Scholar · View at PubMed · View at Scopus
69. J. S. Leong, S. G. Jantzen, K. R. von Schalburg et al., “Salmo salar and Esox lucius full-length cDNA sequences reveal changes in evolutionary pressures on a post-tetraploidization genome,” BMC Genomics, vol. 11, no. 1, article 279, 2010. View at Publisher · View at Google Scholar · View at PubMed
70. J. Piessevaux, D. Lavens, T. Montoye et al., “Functional cross-modulation between SOCS proteins can stimulate cytokine signaling,” Journal of Biological Chemistry, vol. 281, no. 44, pp. 32953–32966, 2006. View at Publisher · View at Google Scholar · View at PubMed · View at Scopus
71. O. Pleguezuelos, J. Zou, C. Cunningham, and C. J. Secombes, “Cloning, sequencing, and analysis of expression of a second IL-1β gene in rainbow trout (Oncorhynchus mykiss),” Immunogenetics, vol. 51, no. 12, pp. 1002–1011, 2000. View at Publisher · View at Google Scholar · View at Scopus
72. K. J. Laing, T. Wang, J. Zou et al., “Cloning and expression analysis of rainbow trout Oncorhynchus mykiss tumour necrosis factor-α,” European Journal of Biochemistry, vol. 268, no. 5, pp. 1315–1322, 2001. View at Publisher · View at Google Scholar · View at Scopus
73. J. Zou, T. Wang, I. Hirono et al., “Differential expression of two tumor necrosis factor genes in rainbow trout, Oncorhynchus mykiss,” Developmental and Comparative Immunology, vol. 26, no. 2, pp. 161–172, 2002. View at Publisher · View at Google Scholar · View at Scopus
74. A. Pezet, H. Favre, P. A. Kelly, and M. Edery, “Inhibition and restoration of prolactin signal transduction by suppressors of cytokine signaling,” Journal of Biological Chemistry, vol. 274, no. 35, pp. 24497–24502, 1999. View at Publisher · View at Google Scholar · View at Scopus
75. J. Piessevaux, D. Lavens, F. Peelman, and J. Tavernier, “The many faces of the SOCS box,” Cytokine and Growth Factor Reviews, vol. 19, no. 5-6, pp. 371–381, 2008. View at Publisher · View at Google Scholar · View at PubMed · View at Scopus
76. S. Ilangumaran and R. Rottapel, “Regulation of IFNγ-induced MHC class-II expression by suppressor of cytokine signaling 1 (SOCS1),” Modern Aspects of Immunobiology, vol. 16, pp. 35–39, 2005. View at Scopus
77. Z. Chen, A. Laurence, Y. Kanno et al., “Selective regulatory function of Socs3 in the formation of IL-17-secreting T cells,” Proceedings of the National Academy of Sciences of the United States of America, vol. 103, no. 21, pp. 8137–8142, 2006. View at Publisher · View at Google Scholar · View at PubMed · View at Scopus